A Taxonomic and Ecological Study of Periphytic Cyanobacteria in Kaituna River and Its Tributaries, Banks Peninsula, New Zealand
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A Taxonomic and Ecological Study of Periphytic Cyanobacteria in Kaituna River and Its Tributaries, Banks Peninsula, New Zealand A thesis submitted in partial fulfilment of the requirements for the degree of Doctor of Philosophy at the University of Canterbury by Faradina Merican bt. Mohd. Sidik Merican School of Biological Sciences University of Canterbury 2013 For Eiman and Dania 2 Acknowledgements First and foremost, my utmost gratitude to my supervisor, Dr. Paul Broady for his valuable guidance, scholarly inputs, consistent encouragement and unconditional support throughout my research work. Paul is a person with an amicable and positive disposition, whose sincerity I will never forget. He is an excellent role model to me as a junior member of academia. Special thanks to Dr. Phil Novis, Landcare Research Lincoln for introducing me to molecular genetics, providing practical advice and commenting on the manuscript. Phil consistently extended his research expertise throughout my work and I owe a lot to him for this achievement. Helpful advice, technical assistance and support were provided by, Reijel Gardiner, John O’Brien, Craig Galilee, Graeme Young, Chriss and Vicki Parr, Roger Lough, Adrian Meredith and Dan Clark. I thank all of them for their contributions. Thanks are extended to Kristy Hogsden and Hamish Greig who have been very kind to extend their help and support in statistical data analysis. To my friends Yatt, Ain, Kak Zita, Ummu, Shaz, Yanti, Salwa, Lan, Mizi, Niza, Abg. Don, Kak Su, K Mas, Abishek and Pavla, a heartfelt thanks. To my kiwi best friend and research companion, Francine Harland, thank you for everything. All these people have provided support and friendship that I will treasure forever The thesis would not have come to a successful completion without the scholarship from Universiti Sains Malaysia and the Ministry of Higher Education Malaysia to support my study. Special thanks to Dean of School of Biological Sciences, Prof Abu Hassan for support and encouragement. To my parents, Mohamed Sidik Merican and Laili Zainudin, and my beautiful family, the biggest thanks of all for all the years of love that provided me with inspiration and was my driving force. To my sweethearts, Eiman Danial and Dania Imani, thank you for all the cuddles and kisses that kept mummy going. Mummy loves you both dearly. Finally, I would like to dedicate this work to my beloved Grandma, Fatimah Nordin who left us too soon. I hope that this work makes you proud. 8 Abstract Most of the detailed studies on periphyton in New Zealand rivers and streams have focused on diatoms. Despite the recent rise of interest in potentially toxic cyanobacterial mats, knowledge of the diversity and ecology of these and other macroscopic growth forms is incomplete. A taxonomic survey was made on periphytic cyanobacteria at 100 locations along Kaituna River and a 1st to 3rd order tributary stream. Samples were taken from runs, riffles and pools in shaded and unshaded locations and from varied substrata from January to December 2011. Descriptions were made of all macroscopic growths. Fifty-six morphospecies were identified of which 29 are new records for New Zealand. Crust components were the most diverse with 23 morphospecies followed by mats (16), gelatinous colonies (5) and epiphytes (7). Five appeared only after growth in enrichment cultures. Twelve morphospecies were isolated into cultures for use in polyphasic assessment. In 16S rDNA phylogenies, Placoma regulare and Heteroleibleinia fontana did not cluster with other members of their traditional families. Nostoc sp. 2 was positioned distant from other Nostoc strains. Comparison of 16S – 23S rRNA internal transcribed spacer compositions for seven mat-forming oscillatorialean morphospecies confirmed their recognition as distinct morphospecies. Amplified fragment length polymorphisms were used to investigate genetic diversity of Nostoc verrucosum in relation to dispersal. This indicated that local dispersal is dominant while cross-catchment dispersal is probably infrequent. Light intensity, substratum type and water conductivity were significant factors influencing spatial patterns of distribution. Higher diversity of crusts, mats and gelatinous colonies was recorded in unshaded locations. Mats and gelatinous colonies were most diverse in Kaituna River and crusts in second to third order streams. Morphospecies in water with high conductivity were restricted to those locations. Spates had a major effect on temporal distribution. An increase in frequency and intensity of spates in spring and winter resulted in greater reduction in cover. Smaller spates caused partial removal followed by rapid regrowth within a week. Major spates caused complete removal of visible cover with re-colonization occurring within three to four weeks. This study has provided a first detailed account of cyanobacterial diversity and ecology in a New Zealand catchment. It provides a basis for long-term monitoring at this site of the effects of changes in climate and in human activities in the catchment. 9 Chapter 1 General introduction 1.1 The context of this study The distribution and diversity of cyanobacteria have not been intensively studied in New Zealand flowing waters (Broady and Merican, 2012). They have been reported as the least common component of stream periphyton after diatoms and green algae (Biggs, 2000b). In many studies, just prominent taxa have been identified to only generic level (Biggs et al., 1998; Biggs and Kilroy, 2000; Biggs and Smith, 2002; Jaarsma et al., 1998). More recently, benthic mat-forming cyanobacteria capable of toxin production have been widely studied (Hamill, 2001; Heath et al., 2010; Wood et al., 2007; 2008; 2010; 2012; Smith, 2010; 2012; Smith et al., 2011). Other important growth forms have received scant attention. These include crusts, tufts, gelatinous colonies and epiphytic communities. Under–sampling and taxonomic uncertainties have resulted in lack of knowledge of the diversity of periphytic cyanobacteria and environmental factors governing their distribution in New Zealand. Previous floristic and ecological studies of periphyton have not specifically focused on cyanobacteria. This has hampered discovery of new taxa that could produce toxins or useful bioactive compounds (Broady and Merican, 2012) and has restricted their use as indicators of water quality. A full knowledge of New Zealand’s biodiversity requires a complete inventory of these widespread microorganisms. The majority of taxa recorded in New Zealand are well-known and cosmopolitan morphospecies while poorly known and possibly endemic species have likely been overlooked (Broady and Merican, 2012). The description of a new species of a colony-forming cyanobacterium, Placoma regulare Broady & Ingerfeld, that is widespread in South Island streams (Broady and Ingerfeld, 1991), suggests that others await discovery. Environmental factors affecting growth and distribution of cyanobacterial periphyton in particular are largely unknown despite numerous detailed studies on New Zealand stream periphyton in general (Biggs, 1990; Biggs and Price, 1987; Biggs et al., 1998; Biggs and Smith, 2002; Jowett and Biggs, 1997; Suren et al., 2003). The ability of cyanobacteria to detect and respond to a variable environment is well-known, e.g. to a wide range in light intensity and spectral quality (Montgomery, 2007; Mullineaux, 2001), periods of desiccations (Garcia-Pichel and Pringault, 2001; Hill et al., 1997), nutrient limitations (Mateo et al., 2006; Whitton, 2008), and grazing pressure (Jang et al., 2007; Yang et al., 2008). Distribution and diversity of cyanobacteria are potentially reliable indicators of water quality in streams and rivers impacted by anthropogenic environmental change 10 Chapter 1 Introduction (Perona et al., 1998; Douterelo et al., 2004; Perona and Mateo, 2006; Loza et al., 2013a). Additionally, morphological changes observed in some morphospecies are also an effect of environmental change, especially nutrient availability (Whitton, 2008). An increase in knowledge of their responses would enhance the use of cyanobacterial assemblages in monitoring the health of New Zealand flowing water ecosystems. 1.2 Project aims and an overview of the structure of this thesis ‘What grows where?’ is the primary question addressed by this research. This study has intensively examined the diversity and distribution patterns of periphytic cyanobacteria in first to fourth order flowing waters of a single catchment on Banks Peninsula, Canterbury, New Zealand. Descriptions were compiled for the diversity of macroscopic growths of cyanobacteria in different microhabitats throughout the study site. Over one year, an assessment was made of the effects of environmental variables on broad and small scale distribution patterns. Chapter 2 describes the study site and features of the catchment and climate. The results are presented in Chapters 3 - 5, each of which provides an introduction and discussion. Chapter 3 describes the diversity of morphospecies of cyanobacteria in the study site. The results presented here respond to the ‘What’ question by addressing morphospecies diversity using descriptions of macroscopic growths and light microscopy of field-collected specimens and, in some cases, isolates studied in culture. Descriptions include photomicrographs and line illustrations. Chapter 4 reports on the molecular phylogenetics of selected cyanobacteria. This responds to the need for polyphasic study in order to fully characterise